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Taphrina Seene Mõju Kase Ksüleemi Morfoloogiale Ja Hüdraulilisele

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Taphrina Seene Mõju Kase Ksüleemi Morfoloogiale Ja Hüdraulilisele EESTI MAAÜLIKOOL Põllumajandus- ja keskkonnainstituut Oles Kapalo TAPHRINA SEENE MÕJU KASE KSÜLEEMI MORFOLOOGIALE JA HÜDRAULILISELE JUHTIVUSELE EFFECT OF FUNGUS TAPHRINA INFECTION ON BIRCH (BETULA PENDULA) XYLEM MORPHOLOGY AND HYDRAULIC CONDUCTION Magistritöö Vee ja maismaa ökosüsteemide rakendusbioloogia õppekava Juhendajad: dotsent Tiina Tosens, PhD Liisa Kübarsepp, MSc Tartu 2018 Eesti Maaülikool Magistritöö lühikokkuvõte Kreutzwaldi 1, Tartu 51014 Autor: Oles Kapalo Õppekava: Vee ja maismaa ökosüsteemide rakendusbioloogia õppekava Pealkiri: Taphrina seene mõju kase ksüleemi morfoloogiale ja hüdraulilisele juhtivusele Lehekülgi: 42 Jooniseid: 4 Tabeleid: 0 Lisasid:0 Osakond: Taimefüsioloogia Uurimisvaldkond: B390 Juhendaja(d): dotsent Tiina Tosens, PhD, Liisa Kübarsepp, MSc Kaitsmiskoht ja -aasta: Tartu, 2018 Käesoleva magistritöö eesmärgiks on uurida Taphrina seene avaldatavat mõju kase ksüleemi morfoloogiale ja hüdraulilisele juhtivusele. Taphrina seen on kottseente hõimkonda kuuluv parasiitseen, hõimkonna tuntuim seen on Taphrina deformans – virsikupuu lehti deformeeriv parasiitseen, mida on kõige rohkem uuritud, kuna see tekitab majanduslikku kahju. Antud töös keskendutakse kaske kahjustavale parasiitseene liigile Taphrina betula (kaseluudik), mis tekitab puuokstele visuaalselt nähtavaid kahjustusi – tuulepesi. Eesmärgi saavutamiseks koguti proove kahest kohast: Tähtvere pargist Tartu linnas ja linna külje all Ihastes asuvalt Anne looduskaitsealalt. Töö käigus uuriti ksüleemitorude suurust ja tihedust, erinevusi mõõdeti fotodelt mikroskoobi all. Selgus, et kontrollproovidel oli ksüleemitorude diameeter suurem kui nakatunud proovidel, seda 9/13, ka oli nakatunud proovidel ksüleemitorude tihedus suurem kui nakatumata proovidel. Töö tulemustes jõuame järeldusele, et Taphrina seenparasiit avaldab mõju kase puidule, mõjutades selle ksüleemitorude suurust ja tihedust. Sellest tulenevalt võib järeldada, et taime hüdrauliline juhtivus on samuti mõjutatud, kuna see sõltub ksüleemitorudest ning nende morfoloogiast, siiski hüdraulilise juhtivuse aspekt vajab tulevikus täiendavat uurimist. parasiitseen, arukask, tuulepesad 2 Estonian University of Life Sciences Abstract of Master’s Thesis Kreutzwaldi 1, Tartu 51014 Author: Oles Kapalo Speciality: Applied Biology of Aquatic and Terrestrial Ecosystems Title: Effect of fungus Taphrina infection on birch (Betula pendula) xylem morphology and hydraulic conduction Pages: 42 Figures: 4 Tables: 0 Appendixes: 0 Department: Plant physiology Field of research: B390 Supervisors: docent Tiina Tosens, PhD, Liisa Kübarsepp, MSc Place and date: Tartu 2018 The aim of this Master’s thesis was to research the influance of the Taphrina fungi on the morphology and hydraulic conductivity of European white birch (Betula pendula). The Taphrina is a fungal genus within the Ascomycota phylum, the most famous species of the division is Taphrina deformans which is infamous of infecting peach trees and is most studied because of economic damage it causes. This thesis focuses on the species Taphrina betulina, which causes visible deformity in a woody plant known as Witch’s brooms. To achieve the goal, samples were collected from two places: Tähtvere Park in Tartu and the Anne Nature Reserv located in Ihaste near the City Tartu. During the study, the size and density of xylem were examined. Differences were measured from photos made by microscope. It turned out that the control samples had a larger xylem diameeter, 9 of 13 samples had larger diameeter, also infected samples had higher xylem density than non- infected samples. In the result of this thesis, we concluded that Taphrina betulina fungus affects the birch wood, affecting the size and density of its xylem. Consequently, it can be concluded that the hydraulic conductivity of the plant is also affected, since it depends on the xylem morphology, however, the aspect of hydraulic conductivity needs further investigation. fungi, birch, Witch’s brooms 3 SISUKORD SISSEJUHATUS ................................................................................................................... 5 1. FOTOSÜNTEES ............................................................................................................... 6 1.1 Fotosünteesi seos produktiivsusega ............................................................................. 7 2. VEEKASUTUSE EFEKTIIVSUS .................................................................................... 9 3. LEHTEDE VEEPOTENTSIAAL ................................................................................... 12 4. VEE LIIKUMINE JUURTEST LEHTEDESSE ............................................................. 15 5. KSÜLEEM ...................................................................................................................... 16 5.1 Kuidas ksüleemi anatoomia varieerub keskkonnastressi tingimustes ....................... 17 6. TAPHRINA SEEN .......................................................................................................... 19 6.1 Kuidas patogeen ründab taime? ................................................................................. 21 8. TULEMUSED ................................................................................................................. 27 9. ARUTELU ...................................................................................................................... 32 KOKKUVÕTE .................................................................................................................... 36 SUMMARY ........................................................................................................................ 37 KASUTATUD KIRJANDUS ............................................................................................. 38 4 SISSEJUHATUS Parasiitseen Taphrina on levinud kogu maailmas, selle levik ulatub peremeestaime leviku areaali piirideni ja on liigist sõltuv (Ludwig-Müller 2015, Каратыгин 2002). Patogeen mõjutab saagikust sellistel taimedel nagu virsik, kurkum, ploom, pirn. Viimaste aastakümnendite jooksul on paljud erinevad puuliigid, sealhulgas ka Põhja- Euroopas kasvavad liigid, olnud invaseeritud erinevate patogeenidega. Seda protsessi on ettevaatlikult seostatud kliima soojenemisega, siiski ei ole enamus teadlasi valmis seostama sellist juhuse kokkusattumust kliimamuutusega. Anda teaduslikku tõendust, mis kinnitaks kliimamuutust kui faktorit, mis põhjustab puude intensiivset koloniseerimist patogeenidega, on äärmiselt keeruline, ning kliimamuutuse ja seenpatogeenide levikut ei tohi tähelepanuta jätta (Hanso, Drenkhan 2013). Võttes arvesse kliima soojenemist ja sellega kaasnevaid keskkonnamuutusi kliimavööndites, kus on omane teistsugune kliima, loob niiske ja soe keskkond ideaalsed tingimused patogeenide levikuks laiuskraadidele, kuhu nende varasem levimine oli keerukam. Taimed puutuvad kokku nii soodsate kui ka ebasoodsate biootiliste teguritega, mis määravad nende ellujäämist taimekoosluses. Stressist mõjutatuna peavad taimed kulutama füsioloogilistes ja biokeemilistes mehhanismides rohkem energiat, et raskendatud keskkonnaoludes ellu jääda (Mandre 2002). Arukask (Betula pendula) on parasvöötmes üks kõige tuntumaid ja levinumaid puuliike, ning kiiresti kasvavava pioneerliigina leidub seda ka raiesmikel, põlendikel ja mahajäetud karjäärides. Samuti on kasel arvestatav väärtus paberitööstuses ja Eestis on suur turumajanduslik huvi selle puidu ekspordi vastu (Tullus, Vares 2001: 9). Käesoleva töö eesmärk oli uurida Taphrina seenpatogeeni mõju arukase ksüleemi morfoloogiale ja veejuhtivusele ning sellega kaasnevat mõju puu produktsioonile. Kontrolliti järgnevaid hüpoteese: (i) Taphirna’ga nakatunud kaseokstel on väiksema pindalaga ksüleemitorud ja seetõttu on taime hüdrauliline juhtivus, fotosünteesi võimekus ja lehe veekasutusefektiivsus väiksem, (ii) Taphrina ei avalda ksüleemile, fotosünteesi võimekusele ja lehe veekasutusefektiivsusele olulist mõju. 5 1. FOTOSÜNTEES Fotosüntees on planeedi biosfääris ainus protsess, mille käigus moodustub hapnik, samuti kaasneb fotosünteesiga orgaanilise saaduse kasv (Miidla 1984). Valgus neelatud fotosünteesi pigmentidena rohelistes taimerakkudes on kasutatud, et võimendada CO2 konverteerimist lihtsateks suhkruteks. Saadud suhkrud kasutatakse ära veelgi keerukamate orgaaniliste molekulide sünteesimisel. Need saadused (st fotosünteesi orgaanilised süsinikusaadused) taimorganism siis kas ladustab lehes või transpordib organismis sinna, kuhu peab vajalikuks (Skillman 2011). Fotosünteesi peetakse energia ja materjali allikaks rakkude metabolismis ja taime kasvus (Skillman 2011). Päikesekiirgus on taimede poolt absorbeeritud ja muudetud biomassiks. Biomassi toodangus on olulisteks faktoriteks organismi kloroplastide jõudlus, päikesevalguse peegeldumine lehtedelt, transpiratsioon, valgushingamine e fotorespiratsioon ja taime metabolism (Silva jt 2015). Fotosünteesi põhiline valem on CO2 + H2O + valgus → süsivesikud + O2 (Komissarov 2014). Atmosfääri gaasilise stabiilsuse tagamisel on fotosünteesil tähtis roll, kuna see tagab CO2 sidumise ja O2 eraldumise (Miidla 1984). Siinkohal tuleb mainida, et kõik elus substantsid koosnevad molekulidest, mis omakorda vormivad taime. Fotosünteesi näol on tegemist reaktsioonidega elektronide ja prootonite tasandil. Antud juhul on väiksema tasandi osakesed footonid, tinglikult võib päikesel toimuvaid tuumareaktsioone vaadelda kui fotosünteesi protsessi algust. Vaatamata fotosünteesi komplekssusele on võimalik määrata kaht peamist protsessi etappi:
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